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Wood and Bark Anatomy of Ranunculaceae (Including Hydrastis) and Glaucidiaceae Sherwin Carlquist Santa Barbara Botanic Garden

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Wood and Bark Anatomy of Ranunculaceae (Including Hydrastis) and Glaucidiaceae Sherwin Carlquist Santa Barbara Botanic Garden Aliso: A Journal of Systematic and Evolutionary Botany Volume 14 | Issue 2 Article 2 1995 Wood and Bark Anatomy of Ranunculaceae (Including Hydrastis) and Glaucidiaceae Sherwin Carlquist Santa Barbara Botanic Garden Follow this and additional works at: http://scholarship.claremont.edu/aliso Part of the Botany Commons Recommended Citation Carlquist, Sherwin (1995) "Wood and Bark Anatomy of Ranunculaceae (Including Hydrastis) and Glaucidiaceae," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 14: Iss. 2, Article 2. Available at: http://scholarship.claremont.edu/aliso/vol14/iss2/2 Aliso, 14(2), pp. 65-84 © 1995, by The Rancho Santa Ana Botanic Garden, Claremont, CA 91711-3157 WOOD AND BARK ANATOMY OF RANUNCULACEAE (INCLUDING HYDRASTIS) AND GLAUCIDIACEAE SHERWIN CARLQUIST Santa Barbara Botanic Garden 1212 Mission Canyon Road Santa Barbara, California 931051 ABSTRACT Wood anatomy of 14 species of Clematis and one species each of Delphinium, Helleborus, Thal­ ictrum, and Xanthorhiza (Ranunculaceae) is compared to that of Glaucidium palma tum (Glaucidiaceae) and Hydrastis canadensis (Ranunculaceae, or Hydrastidaceae of some authors). Clematis wood has features typical of wood of vines and lianas: wide (earlywood) vessels, abundant axial parenchyma (earlywood, some species), high vessel density, low proportion of fibrous tissue in wood, wide rays composed of thin-walled cells, and abrupt origin of multiseriate rays. Superimposed on these features are expressions indicative of xeromorphy in the species of cold or dry areas: numerous narrow late­ wood vessels, presence of vasicentric tracheids, shorter vessel elements, and strongly marked growth rings. Wood of Xanthorhiza is like that of a (small) shrub. Wood of Delphinium, Helleborus, and Thalictrum is characteristic of herbs that become woodier: limited amounts of secondary xylem, par­ enchymatization of wood, partial conversion of ray areas to libriform fibers (partial raylessness). Wood of Ranunculaceae other than Clematis has numerous narrow vessels, probably an adaptation to cold more than drought. Glaucidium has occasional scalariform perforation plates like those almost uni­ versally present in Paeonia, but the two genera differ strongly in other wood features. Wood of Hydrastis (scalariform perforation plates occasional in primary xylem, rare in secondary xylem) ac­ cords with the idea that Hydrastis is a lineage that separated from the base of Ranunculaceae. Features that ally Ranunculaceae with other families of Ranunculiflorae include presence of wide multiseriate rays (without accompanying uniseriate rays), vessel restriction patterns, and storied structure. Wood­ iness in Ranunculaceae is probably secondary, at least to some extent; the woodiness of Xanthorhiza, thought by some to represent a primitive genus in the family, could be either primary or secondary. Key words: Berberidales, Glaucidium, Hydrastis, Paeonia, Papaverales, Ranunculaceae, Ranunculi- florae, storied wood structure, vessel restriction patterns. INTRODUCTION omy of these families may help understand the inter­ relationships of these taxa. The present essay continues Phylogenetically, Ranunculaceae have been of in­ a series begun with studies of Lardizabalaceae (Carl­ terest because they have occupied a near-basal position quist 1984) and Papaveraceae (Carlquist and Zona in many systems. Most recently, the cladograms of 1988; Carlquist et al. 1994). Chase et al. (1993) follow this pattern on the basis of Despite the fact that Clematis contains about 250 rbcL data. Currently, Ranunculaceae are also of inter­ species of woody vines (Willis 1973), only a few spe­ est in relation to the paleoherb hypothesis (Taylor and cies have been investigated with respect to wood anat­ Hickey 1992). Anatomy of wood potentially offers omy. Accounts have been offered by Schmidt (1941), character expressions that illuminate these concepts. Metcalfe and Chalk (1950), Greguss (1959), Grosser Because Ranunculaceae contain a few woody genera (1977), Schweingruber (1978, 1992), and Sieber and as well as many herbaceous genera, it is a key family Kucera (1980). However, these accounts are incom­ in understanding primary versus secondary woodiness, plete, contradictory, and offer some inaccuracies. a chief concern of the paleoherb hypothesis. Wood anatomy of two species of Ranunculaceae in Ranunculiftorae (a superorder that equates with the genera other than Clematis has been briefly described orders Berberidales, Papaverales, Ranunculales, or by Schweingruber (1992), underlining the lack of Rhoeadales of earlier authors) consists of five main wood data for the family. In order to compare wood families: Berberidaceae, Lardizabalaceae, Menisper­ anatomy of Ranunculaceae to that of other Ranuncu­ maceae, Papaveraceae, and Ranunculaceae, as well as liftorae, a more nearly complete and accurate under­ some smaller satellite families. Studies on wood anat- standing of wood in the family than currently at hand is required. In this connection, information is needed 1 Address for correspondence with author: 4539 Via Huerto, Santa on wood of two genera traditionally included in Ran­ Barbara, CA 93110. This study was begun at the Rancho Santa Ana unculaceae (e.g., Prantl 1891): Hydrastis and Glauci­ Botanic Garden, 1500 N. College Ave., Claremont, CA 91711. dium. Recently excluded from Ranunculaceae as 66 ALISO monogenetic families, Glaucidium and Hydrastis have rhosa, and Glaucidium palmatum (see Table 1); por­ been the objects of research, chiefly on embryology tions of these three were preserved in 50% aqueous and floral anatomy (Tamura 1972; Tobe 1981; To be ethyl alcohol. Wood of genera other than Clematis, and Keating 1985). Keener (1993) has offered a sum­ Glaucidium, and Thalictrum were derived from spec­ mary of work to date on Hydrastis, and concludes that imens in the herbarium of the Rancho Santa Ana Bo­ it can be regarded as a subfamily within Ranuncula­ tanic Garden. Some of the wood samples of Clematis ceae. There is now general agreement that Paeonia were obtained from xylaria (see Table 1), and for these should be excluded from Ranunculaceae as Paeoni­ I am especially indebted to Dr. Regis Miller of the aceae and perhaps placed near the families of Ranun­ Forest Products Laboratory (MADw, SJRw) and to Dr. culiflorae (e.g., Thorne 1992), although not within the Roger Dechamps of the Musee Royale de l' Afrique group of families cited above as Ranunculiflorae. Centrale, Tervuren, Belgium (Tw). The woods of Wood anatomy may prove significant in comparing Clematis documented by my collection numbers (Ta­ Paeonia with Glaucidiaceae, Hydrastidaceae, and Ran­ ble 1) are all from cultivated specimens; herbarium unculaceae. specimens to document my collections are deposited Wood anatomy of Glaucidiaceae and Ranunculaceae at the Rancho Santa Ana Botanic Garden. I am grateful is worthy of examination with respect to habits and to the Municipal Botanic Garden, Dunedin, New Zea­ habitats. The woody vine Clematis ranges from upland land, and to the Rancho Santa Ana Botanic Garden for tropical to temperate alpine sites. The "woody herb" wood from cultivated specimens included in this study. (herb, with various degrees of secondary xylem pro­ The stem of Thalictrum polycarpum was collected duction in stems near the ground or beneath the ground from cultivated plants in the Santa Barbara Botanic surface) habits of some Ranunculaceae should be Garden. taken into account in studies of wood anatomy. Of the The localities of noncultivated specimens are as fol­ genera included in this study, only Xanthorhiza qual­ lows: C. dioscoriifolia, ruderal, New York City; C. ifies as a shrub or subshrub, an understory element in haenkeana, Ecuador; C. iringaensis, Shaba, Central moist temperate forests. Some species of the North Africa; C. ligusticifolia, dry riverbank near Yakima, Temperate genus Delphinium have a short, upright, Washington; C. paucifiora, Santa Ana Mountains, Cal­ woody caudex. Helleborus (Mediterranean region to ifornia; C. pickeringii, Fiji; Delphinium elatum, 83 the Caucasus Mountains) can be a subshrub with mod­ miles from Novosibirsk on highway to Bernaul, Si­ erately woody stems (H. foetidus L.), but most species, beria, Russia; Glaucidium palmatum, Mt. Shirano near such as H. viridis studied here, have rhizomes with Lake Chuzenji, Japan; Helleborus viridis, Montelupo, only a little secondary xylem. Hydrastis (two species Italy; Hydrastis canadensis, Neversink, Berks Co., of moist temperate forests: H. canadensis and H. je­ Pennsylvania; Xanthorhiza apiifolia, MacDowell Co., zoensis Sieb. & Zucc.) has succulent underground rhi­ North Carolina. The identity of the specimen cited here zomes. Glaucidium has a habit similar to that of Hy­ as C. haenkeana was determined with the aid of com­ drastis, but with more condensed rhizomes; the single ments from Dr. Frederick Essig and Dr. Nancy Mo­ species is confined to montane northern Japan (Prantl reno. 1891). Genera with minimal secondary xylem have Dried wood samples of both Clematis and the re­ been excluded from this study. maining genera (incorporating bark portions) provided Are Ranunculaceae primarily or secondarily woody, difficulties in sectioning because of wideness of ves­ or have both phyletic curricula occurred within the sels and (in bark) contrast between softness of phloem family? Wood anatomy examined in this context can and hardness of fibers. These problems were solved by cast light not only on this family but on others (such using a technique involving
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